Magnetoreception in microorganisms and fungi · fungi, and try to show how the variegated empirical...

DOI: 10.2478/s11535-007-0032-zReview article

CEJB 2(4) 2007 597–659

Magnetoreception in microorganisms and fungi

Alexander Pazur1∗, Christine Schimek2, Paul Galland3†

1 Department of Biology ILudwig-Maximilian University Munchen,

D-80638 Munchen, Germany2 Department of General Microbiology and Microbial Genetics

Friedrich-Schiller-University JenaD-07743 Jena, Germany

3 Faculty of BiologyPhilipps-University MarburgD-35032 Marburg, Germany

Received 14 May 2007; accepted 09 July 2007

Abstract: The ability to respond to magnetic fields is ubiquitous among the five kingdoms of organisms.Apart from the mechanisms that are at work in bacterial magnetotaxis, none of the innumerablemagnetobiological effects are as yet completely understood in terms of their underlying physical principles.Physical theories on magnetoreception, which draw on classical electrodynamics as well as on quantumelectrodynamics, have greatly advanced during the past twenty years, and provide a basis for biologicalexperimentation. This review places major emphasis on theories, and magnetobiological effects thatoccur in response to weak and moderate magnetic fields, and that are not related to magnetotaxis andmagnetosomes. While knowledge relating to bacterial magnetotaxis has advanced considerably duringthe past 27 years, the biology of other magnetic effects has remained largely on a phenomenological level,a fact that is partly due to a lack of model organisms and model responses; and in great part also to thecircumstance that the biological community at large takes little notice of the field, and in particular ofthe available physical theories. We review the known magnetobiological effects for bacteria, protists andfungi, and try to show how the variegated empirical material could be approached in the framework ofthe available physical models.c© Versita Warsaw and Springer-Verlag Berlin Heidelberg. All rights reserved.

Keywords: magnetic field, magnetoreception, ion-cyclotron resonance, magnetosomes, quantumcoherence, radical-pair mechanism, ecology, climate change

∗ E-mail: [email protected]† E-mail: [email protected]

598 A. Pazur et al. / Central European Journal of Biology 2(4) 2007 597–659

Abbreviations

B magnetic flux density (magnetic induction)

BAC alternating magnetic field (generated by alternating current)

BDC static magnetic field (generated by directed current)

CD coherent domain

ELF extremely low frequency (i.e. magnetic field, ∼3-300 Hz)

EMF electromagnetic field

H magnetic field strength

IIM ion interference mechanism

ISC intersystem crossing

ICR ion cyclotron resonance

IPR ion parametric resonance

LF low frequency (i.e. magnetic field)

MF magnetic field

1 Introduction

The earliest studies on the influence of electromagnetism on organisms date back to the

late 19th century, probably beginning in St. Petersburg [1]. A larger, more general interest

arose only some decades later, coinciding with worldwide electrification and telecommuni-

cation. Although microorganisms play a major role in the global ecosystem, the number

of publications covering magnetoreception in fungi, protists and non-magnetotactic bac-

teria is small compared to similar reports on humans and animals [1–6]; and is perhaps

comparable to the state of knowledge in plants [7]. The magnetoorientation of mag-

netotactic bacteria [8], as well as that of migrating birds and insects, belongs to the

best understood and most intensely studied phenomena of magnetoreception [6, 9, 10].

Recently Ritz et al. [11, 12] suggested a light-driven, radical-pair mechanism for the mag-

netoreception of birds mediated by cryptochrome [13, 14]. There is evidence that even

plant cryptochromes are involved in the magnetoreception of Arabidopsis [15]. Bacterial

magnetotaxis is based on the magnetoorientation of magnetite crystals; thus representing

the only magnetoreception mechanism completely elucidated up to now [8, 16–18].

The two central questions in this context: (i) whether or not microorganisms are

able to perceive geomagnetic fields, and (ii) whether or not magnetoreception is an es-

sential and vital environmental factor for survival, have remained largely unanswered,

even though magnetoreception must be regarded as an established fact. Furthermore,

we contend that the recent discussion regarding the mechanisms of climate change and

global warming should consider other, non-anthropogenic contributions, e.g. the altered

gas exchange of microorganisms as a consequence of the steadily changing geomagnetic

field.

Despite the numerous magnetobiological effects that have been described in the per-

tinent literature, there is an apparent lack of model organisms, model responses and

genetic approaches; tools that are typical for modern research strategies commonly found

A. Pazur et al. / Central European Journal of Biology 2(4) 2007 597–659 599

in other biological fields. The problem is compounded by the fact that magnetic effects

are observed for a huge range of magnetic flux densities, which cover more than 10 or-

ders of magnitude. To come to grips with such a huge dynamic range, that is similar to

that of human vision, one would expect the study of dose-response relationships to be

of paramount importance. It thus comes as a surprise that there exists only one dose-

response curve for a biomagnetic effect in DC fields [19], and only a limited number of

dose-response studies for AC-fields. Despite the limited information it is nevertheless ap-

parent that magnetobiological dose-response relationships differ drastically from the ones

usually found in physiology, where one typically finds exponential rise or decay curves,

and in some cases optimum curves. Magnetoresponses, in contrast, show certain “win-

dows” of magnetic flux densities, or, in the case of AC-fields, windows of frequencies for

which a response is obtained. Applying a higher field strength may thus not necessarily

guarantee a stronger response. As a consequence, some of the apparent contradictory

results in the magnetism literature could be explained by the fact that different authors

often used different magnetic flux densities that were within or outside these windows.

Research on biological magnetoresponses can be roughly divided into experiments

that employ static magnetic fields (BDC), or alternating fields (BAC), or as in most cases,

a combination of both (BAC+ BDC). The body of literature on AC fields and their

concomitant effects dominates by far that of DC fields. This appears surprising in view

of the fact that DC experiment is required to find out how geomagnetic fields influence

life.

Most experiments with AC fields are done with frequencies near 50 or 60 Hz, i.e.

frequencies akin to that of ubiquitous electric appliances. Much of this type of research

was historically motivated by the wish to find out whether or not our electric environment

influences life, and specifically, human health. Even though this line of research may

not directly contribute to the understanding of how static geomagnetic fields influence

life, it nevertheless represents a powerful technical tool to investigate the involvement of

specific ions in a given biomagnetic response. It had earlier been noticed that AC fields

elicit responses most prominently at the cyclotron resonance frequencies (including their

harmonics and subharmonics) of biologically important ions, in particular Ca2+. This

pattern gives rise to dose-response curves with several minima and maxima. Therefore

it is not difficult to understand why explaining this type of dose-response relationship is

the subject of several theories (ion-cyclotron resonance, ion-parametric resonance, ion-

interference mechanism, coherence mechanism; see below).

One of the reasons that magnetobiological responses frequently meet with reserva-

tions is based on the fact that the energy content of biologically actinic magnetic fields

can be several orders of magnitude below their thermal energy content (kT problem).

We will show how the problem can be addressed within the framework of modern the-

ories. Also the hunt for “the” magnetoreceptor remains presently an unresolved task

(with the exception of magnetite in bacterial magnetotaxis, see below). As function of

fact, it is contested as to whether or not there exists only one type of magnetoreceptor;

the requisite literature rather indicates that in prokaryotic and eukaryotic cells several


magnetosensitive molecules, and physically distinct mechanisms, exist that could mediate

magnetoreception. Because cell membranes do not constitute barriers for magnetic fields,

magnetoreception could in theory occur on many different levels. For example, DNA it-

self, and the transcription and translation machineries, have been proposed as targets.

It is thus noteworthy that even cell-free systems of protein biosynthesis are receptive to

magnetic fields (see below).

2 Magnetic effects on solutes

Magnetic fields can affect organisms not only directly, but also indirectly, by changing

the physical properties of solutes and growth media. For example, a 1 minute magnetic

pretreatment of culture media stimulated the subsequent growth of Escherichia coli in a

geomagnetic field [20]. Magnetically treated water inhibits the germination of the micro-

fungus Alternaria alternata [21], and treatment of nutritional media affects the subse-

quent growth of Saccharomyces fragilis, Brevibacterium and Bacillus mucilaginosus [22].

Such effects are possible because magnetic treatments alters solutes, for example the

formation of calcium carbonate [23], water vaporization [24], ion hydration and resin

absorption [25]. Indeed effects on Ca2+ hydration after short treatment with a weak

magnetic field or pulses, applicable, for example, to organismal growth stimulation, is

reported by Goldsworthy et al. [26]. As these effects were usually achieved with very

strong magnetic fields in the mT to T range, they may not be pertinent for experiments

done in very weak or geomagnetic fields.

3 Bacterial magnetotaxis

3.1 Magnetosomes and their role in magnetoreception

Apart from the general effects of all types of magnetic fields on growth and morphogen-

esis, some organisms have succeeded in employing the directional qualities of magnetic

fields for orientation purposes. Animals using geomagnetic fields for navigation are either

long-distance travellers, such as migratory birds, whales, sharks, turtles and butterflies,

or depend for other reasons on the ability for exact orientation, e.g. honeybees. Clearly,

microorgansims do not fall into a category where magnetoorientation would be expected.

Nonetheless this behaviour, known as magnetotaxis, is the best studied of the magnetore-

sponses [27–29]. This reaction has been globally observed in a number of marine [30, 31]

and freshwater bacteria [32–34], as well as in several types of unicellular eukaryotic mi-

croorganisms. The latter are rarely observed, probably because they are easily overlooked

in samples teeming with bacteria. Due to their overall high fragility and sensitivity, eu-

karyotic laboratory strains usable for detailed analyses have not yet been established; yet

the occurrence of magnetoperception in eukaryotes may be rather widely distributed, as

magnetotactic species have been detected in a number of major groups, such as dinoflag-

ellates, ciliates, cryptophytes [35, 36].


3.2 Magnetotactic microorganisms

Research into magnetoresponsive prokaryotes began with the discovery that certain bac-

teria consistently preferred one geomagnetic pole over the other, and therefore always

swam to the same side of a water droplet on microscopic slides [16]. Magnetotaxis, al-

though biased for fast-swimming organisms, provided a handy tool for the isolation of

several similar species [4, 37–39, 41–45]; and moreover also allows for the enrichment

and studies of unculturable strains and communities [46, 47]. Magnetotactic bacteria are

flagellated chemolithoautotrophs exhibiting various morphotypes; to date cocci, spirilla,

rod-shaped, and vibroid or helical forms have been described [47, 48]. They inhabit the

oxygen-anoxygen transition zone of marine or freshwater sediments or chemically strati-

fied water columns, where they occasionally occur in high cell densities [31, 49] and are

either obligate anaerobes [30, 45] or facultatively anaerobic microaerophils [50–53]. Al-

though likely to be of polyphyletic origin [54], most of the characterized morphotypes

have been grouped into the Proteobacteria, with a distinct subcluster present in its al-

pha subgroup [32, 42, 46, 55, 56] (Table 1). Genome data are available from Magne-

tospirillum magnetotacticum MS-1 (GenBank accession AAAP00000000, Magnetospir-

illum gryphiswaldense MSR-1A [57]; GenBank acc. BX571797) and Magnetospirillum

magneticum AMB-1 (GenBank acc. AP007626). Highly organized aggregates of magne-

totactic cells have also been described from a variety of other locations [28, 58–61].

3.3 Magnetotaxis

Magnetotaxis is defined as movement parallel to the field lines of an external magnetic

field. Nevertheless it is not a taxis in the strictest sense as the organisms are not following

the direction of the magnetic field itself, but rather ultilize the directional information

to support other orientation mechanisms. It was generally assumed that they navigate

along the inclination of the magnetic field lines to locate a suitable environment within an

oxygen (magneto-aerotaxis), or other chemical, gradient; thus reducing search movement

in turbid surroundings to just one dimension - up and down [62]. The key benefit of

magnetotaxis in this process is the enhancement of the bacterium’s ability to detect

oxygen, not an increase in average speed of reaction [29]. Movement along a straight

path allows for earlier detection of an existing oxigen gradient, and thus enhances the

flight from oxygen. One study suggests a role for magnetosome formation in mediating

the response to gravity, as magnetosomes and magnetotaxis were shown to be completely

absent in prolonged microgravity [63]. In magnetotaxis, polar and axial magnetotactic

strains can be discriminated between. Bipolar flagellated cells display axial behavior by

swimming back and forth within a local applied magnetic field. In polar magnetotaxis, the

cells follow a preferential direction and swim away when the local field is reversed [64].

This classification apparently results from cellular morphology, and has no impact on

orientation efficiencies in natural environments. The observation that polar magnetotactic

cells in the southern hemisphere predominantly exhibited a south-seeking behavior in


laboratory tests was taken as support for the importance of the magnetic field line for

magnetotaxis [65]. The recent discovery of seasonally occurring, predominantly south-

seeking polar bacteria, in populations from the northern hemisphere call this explanation

into question. Instead, the oxidation-reduction potential at any given position of a water

column seems to influence the polarity of movement [31, 49].

3.4 Magnetosomes

All magnetotactic cells contain magnetosomes. These organelles consist of a ferrimagnetic

crystal surrounded by a specialized membrane. In prokaryotes, the magnetosome crys-

tals result from the controlled biomineralization of either magnetite (Fe3O4), or greigite

(Fe3S4) [66–69]. Additionally one single strain has been found that contains iron pyrite

(FeS2) [67]. A few morphotypes mineralize magnetite and greigite within the same cell,

and even within the same crystal aggregate [70, 71].

In both magnetite and greigite, crystal structures follow the spinel type, consisting of

two interlocking grid systems with different numbers of grid coordinates (nodes). Mag-

netite, as well as greigite, contains a mixture of two- and three-valent iron, with each

form occupying specific nodes. This leads to the complete extinction of the atomic mag-

netic dipol moments of Fe3+. The magnetic properties, therefore, are solely attributed to

Fe2+. Each morphotype is usually associated with a particular crystalline habit of mag-

netite, whereas greigite crystals of different shapes may occur simultaneously [48, 72–74].

Cuboid, bullet-, tooth- and drop-shaped crystals have been described [45, 73, 75, 76].

Besides eukaryotic microorganisms, magnetite crystals that are similar in appearance

and structure to those of bacteria were also found in animal cells [77]; however no infor-

mation exists on their origin and biosynthesis. Ferrimagnetic crystals interact in excess of

a million times more strongly with magnetic fields than do diamagnetic or paramagnetic

materials. If a ferrimagnetic nanocrystal were fixed to an ion channel - an assumption

that has not been verified yet - it would generate torque in a weak geomagnetic field that

would suffice to alter ion movement across a membrane. Such considerations show that

magnetites hold, at least in theory, the potential to directly influence ion transport [77].

It has also been pointed out that trace amounts of magnetite may be ubiquitous, and

that a single 100-nm magnetite crystal, exposed to a 60 Hz, 0.1 mT magnetic field, could

absorb sufficient energy to supersede several times the thermal background noise [78].

Magnetite particles can have dramatic effects on the dynamics of photogenerated free

radicals [79]. It is thus pertinent to reckon with a modulating effect of magnetites if

present, particularly in context of the radical pair mechanism (see below).

Fossil records of magnetosome crystals date back to the Precambrian time; while sim-

ilar crystals have been detected in 4 billion year-old carbonate blebs of martian meteorite

fragments [80–82]. Although controversely discussed [83, 84], it appears possible that the

martian magnetites are of a biogenic origin. This would also imply that these martian

minerals constitute the oldest fossils on Earth, and at the same time provide evidence for

the possibility of panspermia [85].


The number of magnetosomes within a single cell ranges from a few large structures

in cryptophyte cells [35] to several hundred [73, 86], with 10 − 20 the average number

in magnetotactic spirilla [8, 16, 72]. Crystal sizes range from 35 to 200 nm, which in-

dicates their single-domain status [86–88]. The size of a magnetic domain depends on

the material, and can be roughly calculated. According to such estimations, a domain

of magnetite corresponds to a size between 35 and 75 nm, and in elongated crystals up

to 120 nm [89–91]. As single domain crystals, the magnetosome cystals are especially

susceptible to efficient magnetization and alignment, and they produce stable magnetic

fields. Exceptions have been published by Farina et al. [92] and Spring et al. [56], who

demonstrated that at least two strains isolated from the Itaipu lagoon in Brazil contained

magnetosomes with dimensions up to, and even exceeding, 200 nm, a size that could eas-

ily harbour two magnetic domains. In such large crystals a metastable, single-domain

state is only possible when the crystals are aligned within a chain [93]. The extracellu-

lar formation of single-domain magnetite for biotechnological applications has also been

performed by a biologically-induced, biomineralization process of non-magnetotactic bac-

teria [94, 95]; and by the aerobic fungi Fusarium oxysporum and Verticillium sp. [96].

3.5 Magnetosome organization and synthesis

In some morphotypes magnetosomes form loose aggregates within the cell [60, 97], how-

ever in the majority of strains studied they are arranged in one or more chains spanning

the cytoplasm. The magnetosomes within a given chain are separated by a gap containing

no particulate structures, as observed in transmission electron micrographs [98]. In Mag-

netospirillum species, the single magnetosome chain is usually located close to the inner

membrane [98, 99]. The combination of disposition in chains, and size control, results in

a high magnetic to thermal energy ratio. The total magnetic moment of a magnetosome

chain equals the sum of the individual particle moments [100], and substantially surpasses

thermal noise [73, 101].

Organization into chains implicates the crystals in magnetizing each other, and align-

ing their magnetic dipole moments with each other. These processes start at synthesis,

thus each newly formed magnetosome crystal is influenced by the pre-existing chain. Bio-

logically controlled biomineralization is a highly precise process, and is necessarily subject

to very exacting control. Therefore the organism first creates a matrix, delimiting the

space within which the mineral will grow. The form and size of the nascent crystals

depend on the interactions between organic and inorganic phases, and are influenced by

parameters such as pH, redox conditions, ionic strength, lattice geometry, polarity, stereo-

chemistry and topography. The biomineralization of greigite is less well studied than that

of magnetite. It seems to be less organized, and to require considerably more time [102].

Similar to magnetite biomineralization, it requires several mineralization steps, leading

from the non-magnetic precursors, mackinawite and cubis FeS [103], to the final prod-

uct over a transition period of several days or weeks. During this latter period, iron atoms


are rearranged between adjacent sulfur layers, and some of the iron is lost and likely

deposited as amorphous iron oxide aggregates.

The processes leading towards the formation of magnetite have been reviewed by

Schuler [64, 72, 76, 104–106]. At the onset, a low oxygen potential is likely to be a

regulatory signal for metabolic induction of biomineralization, as in Magnetospirillum

gryphiswaldense, M. magnetotacticum and Magnetospirillum sp. AMB-1. Thus biominer-

alization only occurs at pO2 values below a threshold of 20 mbar [107]. Biomineralization

occurs inside a specialized organella, the magnetosome, providing a scaffold organized by

membrane proteins which ensures the spatial and temporal accuracy of the process. The

scaffold need not be proteinaceous: it can also be thought of as a matrix of amporphous

mineral precursors [108]. Indeed amorphous iron oxide has been found to form a layer sur-

rounding maturing crystal [102]. Magnetosomes are enmeshed in a network of cytoskeletal

filaments [99], and provides a surrounding for the precise coordination of events involved

in magnetite biomineralization [109]. The whole complex consists of several structural

entities: the magnetite crystal, magnetosome membrane, surrounding matrix, and, as

described for Magnetospirillum magnetotacticum MS-1, an interparticle connection [110].

It has been assumed that magnetosomes are invaginations of the cell membrane; indeed

proteins probably involved in such an invagination process have in fact been identified in

the magnetosome membrane [111]. Recently electron cryotomography revealed that the

membrane surrounding magnetite crystals is continuous with the inner membrane [99].

Nevertheless, some questions remain. The process of iron acquisition and biomineraliza-

tion would require a closed compartment. Also, in the electron cryotomography picture

series, the connection of magnetosomes with the inner membrane was only visible for

the innermost structures; the largest magnetosomes seemingly already contained finished

crystals. The small, incomplete magnetosomes at the chain ends were completely inside

the cytoplasm, with no apparent contact with the inner membrane. Moreover, the lipid

and protein composition of the magnetosome membrane differed from all of the other

membrane systems of the cell [112]. If it really does originates from the inner membrane,

then it is at the very least least subject to extensive modifications. The hitherto iden-

tified proteins are apparently involved in iron import, iron conversion and in magnetite

synthesis [57, 113, 114].

These membrane vesicles precede magnetite biomineralization and may exist inde-

pendently [109]. When cells grown under iron limitation are changed to iron sufficiency,

biomineralization occurs simultaneously in many pre-formed vesicles, and from the same

location within each vesicle. In cells with sufficient iron supply, new magnetite crystals

are formed in vesicles at the end of the fully developed magnetosome chain [109]. Usually,

the magnetosome chain is distributed to daugther cells at the point of cell division, and

during cellular growth. However complete de novo synthesis is also possible [71].

As a prerequisite for biomineralization, iron needs to be imported into the cell. This

is a very fast process, for example in iron depleted cells of Magnetospirillum sp. AMB-1

iron uptake is complete within 10 minutes [115], and in Magnetospirillum gryphiswaldense

an increase in magnetite measured as intracellular insoluble iron can also be found within


10 minutes [116]. Indeed a recent microarray analysis of iron-inducible genes demon-

strated the up-regulation of Fe2+ transporters in iron-rich conditions [115].

Soluble Fe(II) may be taken up by the cells by unspecific mechanisms. In these cells,

Fe2+ was found asssociated with the cell envelope, whereas the free Fe3+ was associated

with the magnetosomes [117]. Other strains use Fe(III), and require more complicated

translocation systems. Magnetospirillum magnetotacticum, which has an iron content

approaching 2% of its total dry weight, incorporates iron as Fe3+ [37] using a siderophore

transport system [118]. A ferric iron reductase was purified and characterized by Noguchi

et al. [119] and may well be involved in the subsequent periplasmatic reduction of Fe3+.

Iron oxidation in Magnetospirillum magnetotacticum MS-1 is an aerobic respiratory pro-

cess, and is also necessary for magnetite synthesis [120]. In a generalized scheme, Fe(III)

becomes reduced upon entering the cell. The resulting Fe(II) is then incorporated into

empty magnetosome vesicles, already possessed of its specific protein components. Inside

the magnetosome Fe(II) becomes oxidized again. From this process hydratized Fe(III)-

oxides result, which are dehydratized step by step. Just before the last dehydration step,

at the final position within the crystal, one third of all of the Fe(III) is reduced again to

form Fe(II). The final dehydration step then leads to the product, magnetite [121].

The magnetosome is connected via a membrane protein to the cytoskeleton, and is

thus fixed at a distinct postion within the cell [99, 122]. The chain results from the

magnetosomes being attached to filaments of the actin homologon, MamK [99, 122].

This close contact between magnetosomes and the cell membrane may provide additional

stability to the chain [123].

The genes encoding proteins of the magnetosome membrane, or those otherwise in-

volved in magnetosome formation, are concentrated in several clusters [124–126]. Within

the magnetotoactic bacteria, the homology of individual genes is high, and their positions

within clusters are well conserved. Some of the genes have been identified and are com-

prised of several that would encode proteins with protein-protein interaction domains,

e.g. tetratricopepetide-repeat-proteins. Thus protein complexes may be necessary for

events in the biomineralization process.

Other groups encompass genes specifying proteins involved in: metal transport, other

forms of transport, and those for protein processing, e.g. chaperones or specific proteases.

Within these clusters the gene for the actin homolog is also present. The magnetosome

gene clusters are themselves concentrated on a genomic island, a region of about 130 kb

existing exclusively in magnetotactic bacteria. Deletion of this region leads to the com-

plete loss of magnetosomes and magnetotaxis [57, 124, 126]; yet curiously the functional

magnetosome island alone is insufficient for magnetosome formation. Thus orthologs

present outside of this discrete genetic island are also necessary for this process [126].

Within this region, a conspicuous number of insertion elements, transposases, integrases

and other phage - associated genes abounds [124]. This accumulation, and the high and

retained homology between different strains, hints at the probability that the magneto-

some gene island was acquired via phage-mediated, horizontal gene transfer. The region


was also found to be hypervariable, as spontaneous mutations displaying various phe-

notypes often occur, especially during stationary growth.

Within the magnetosome chains, the individual units attract each other via their im-

manent magnetic forces. This implies that their opposite magnetic poles are facing each

other, and thus their magnetic dipole moments are mutually enhancing each other. Chain

formation is influenced by the shape of the crystals, with rigid chains more easily main-

tained by cubic shapes, as compared to tear-shaped magnetite [123]. In fact, a chain of

magnetosomes corresponds to an equivalently sized bar magnet spanning the whole cell

which generates a permanent magnetic dipole moment [100]. The magnetic field within

a magnetite crystal chain can even be visualized using modern electron microscopical

techniques [91, 127].

The crystal chain is rigid within the cell, and is fixed in position. As a bar magnet,

the magnetic moment is sufficiently large to align with the geomagnetic field. The cell

is drawn with the magnetosome chain, and thus becomes aligned passively and parallel

to geomagnetic field lines [100]. This phenomenon also occurs with dead cells, although

only living ones may move along magnetic field lines. Passive attraction to a magnetic

pole is also possible [128].

4 Zero and weak magnetic fields

To asses the role of magnetic fields in nature, one depends on investigations done at

geomagnetic field strengths (75 μT at the poles - 25 μT at the equator), and, in addition,

also under conditions without a magnetic field (zero field = magnetic vacuum). Apart

from studies on bacterial magnetotaxis, such investigations are, however, extremely rare;

and it remains largely unknown what role geomagnetic fields plays in nature.

Weak static MFs (0 − 110 μT) affect the “anomalous viscosity time dependence” of

E. coli, a parameter that reflects the status of DNA-protein complexes [19]. Interestingly,

dose-response curves for this effect show several minima and maxima. These observations

were explained using the framework of the ion interference mechanism, and were linked

to the dissociation of ion-protein complexes that rotate at a speed of about 18 revolutions

per second. The authors believe that the carrier for the rotating, ion-protein complexes

is DNA [19]. E. coli, Pseudomonas and Enterobacter display, in a zero-magnetic field,

modified resistance to various antibiotics [129, 130]. Surprisingly, even the extremely low

magnetic flux densities generated by the human body can affect bacteria, because E. coli

and Staphylococcus aureus have altered functional activities [131]. For fungi and protists

no studies on the effect of zero magnetic fields are presently available.

Geomagnetic storms can lead to a small increase in geomagnetic fields by some 1−5%.

This increase seems to be sufficient to prolong the photobioluminescence of Photobac-

terium [132]. In the slime mold, Physarum polycephalum, a weak field of 100 μT elicits

a mitotic delay, and decrease of respiration [133]. At a magnetic flux density of 100 μT,

the growth of the phytopathogenic fungi, Alternaria alternata, Curvularia inaequalis and


Fusarium oxysporum, decreased by some 10%. At the same time the MF caused an

increase of conidia formation in A. alternata and C. inaequalis by some 68− 133% [134].

5 Effects on growth and cell division

5.1 DC-fields

Very strong magnetic fields (5.2 − 6.1 T) are able to delay cell death in stationary cul-

tures of Bacillus subtilis [135]. A field of 14.1 T had, however, no substantial effect

on the growth of Shewanella oneidensis, even though several genes were up- or down-

regulated [136]. The latter result shows that growth can be highly inappropriate for

evaluating the magnetosensitivity of an organism.

Moderate static magnetic fields (0.1 − 1 mT) stimulated, both in liquid and solid

media, the growth and metabolism of Pseudomanas fluorescens, Staphylococcus albus

and Aspergillus niger [137]. In contrast, three species of Acanthamoeba responded with

a growth decrease at modest static fields of 71 and 106 mT [138]. A weak static field

(400 μT, i.e. about 8 times the geomagnetic field) elicits, in Saccharomyces cerevisiae,

a 30% inhibition of bud formation [139]. Colonial growth of Alternaria alternata and

Curvularia inaequalis decreased by a mere 10% during exposure to weak magnetic fields

between 0.1 and 1 mT [140]. An inhibition of growth was reported for Anabaena doliolum

for a moderate DC field of 300 mT [141].

5.2 AC-fields

Numerous investigators have reported magnetic effects on development of bacteria, which

includes an increase in mass and / or cell division. Escherichia coli, for example, when

exposed to an AC field (0−22 mT, 16 and 50 Hz), shows a shortened generation time [142].

The dose-response relationships for this effect were complex, they occurred only at certain

flux densities between 0 and 22 mT. AC fields (0.8, 2.5 mT, 0.8 and 1 kHz) and increased

the growth of Bacillus subtilis, as it caused a growth increase and interestingly also a loss

of intercellular cohesion, which is characteristic for cells raised in a geomagnetic field [143].

Whether or not an AC magnetic field exerts an inhibitory or else a stimulatory mode

of action depends in a complex manner on the frequency and the field strength. For

example, Moore [144] observed elevated or even diminished growth rates for Bacillus

subtilis, Candida albicans, Halobacterium, Salmonella typhimurium, and Staphylococci in

dependence of AC frequencies ranging from 0 - 0.3 Hz and magnetic flux densities of

5− 90 mT. In contrast, magnetic square wave signals (0.05− 1 mT, 50 Hz) had no effect

on the growth of E. coli [145]. The viability of Escherichia coli, Leclercia adecarboxylata

and Staphylococcus aureus was negatively affected by prolonged exposures to AC fields

of 10 mT, 50 Hz) [146].

In Paramecium tetraurelia, AC fields (1.8 mT, 72 Hz) caused increased cell division

rates, a response that was Ca2+ specific, and absent in the presence of a Ca2+ blocker.


The magnetic treatment also caused alterations in membrane fluidity [147]. Physarum

polycephalum responds to weak AC fields (0.2 mT, 60, 75 Hz) with a delay in its mitotic

cycle [133, 148], exhibited by an increased mitotic cycle length at 0.2 mT and 75 Hz [149–

151].

The mechanism for magnetotactic effects at ELF-frequencies (e.g. 50, 60, or 75 Hz) is

not completely clear, however energy conversion to heat can likely be ruled out because

of the low induction of living matter. Conversely higher frequency, long wave band fields

(160 mT, 62 kHz) are in fact lethal for E. coli [152]. After an exposure time of 16 h

only a small fraction (10−4 organisms) survive. Under these conditions, the dissipation

to heat is likely not to be increasingly negligible, and, in general, these results are not

comparable with findings for the ELF band in any case.

6 Effects on DNA: mutagenicity, repair, transposition

Weak, static magnetic fields (0−110 μT) affect DNA-protein conformations in E. coli [19].

This analysis represents the only dose-response curve for a static magnetic field. The

peculiarity of this curve stems from the fact that it has three prominent maxima, a

feature that makes it very different from other dose-response curves in nature that often

follow rising or decaying exponential functions. The shape of this curve is explained in

the context of the ion interference mechanism [19].

AC fields (14.6 mT, 60 Hz) have been shown not to cause DNA breaks in a Salmonella

test system [153]. Various strains of Escherichia coli, including DNA-repair mutants,

showed no evidence of increased DNA damage when exposed to very strong magnetic

fields (0.5 and 3 T) [154].

The transposition frequency of Tn10 in E. coli is enhanced by pulsed, square-wave

AC fields [145], but is diminished by sinusoidal AC fields [155]. Increased transposition

activity was also obtained for Tn5, after the exposure of E. coli to AC magnetic fields

(1.2 mT, 50 Hz). Concomitantly, DNA repair was enhanced [156], an event that was

seemingly mediated by the overexpression of DnaK/J [157].

AC fields (0.2 mT, 60 Hz) can increase in Salmonella typhimurium, azide-induced

revertants [158]. The enhanced DNA repair of hydroxylamine-mutagenized plasmid pUC8

occurred in E. coli in AC magnetic fields (0 − 1.2 mT, 50 Hz) via the induction of heat-

shock proteins Hsp 70 and Hsp 40 (DnaK and DnaJ) [156]. Since it is known that DnaK

can upregulate UvrA, it is understandable that magnetic field stress causes improved

DNA repair. An AC magnetic field also (120 μT, 50 Hz) caused a reduction in the

survival of Saccharomyces cerevisiae after UV irradiation, whilst sustaining no effect on

cell cycle kinetics [159].

Most of the effects listed in Tables 2–5 are generally modest, i.e. often amounting

to a response of some 10% to maximally 50%. A notable exception is the response of

E. coli to strong unhomogenous fields (5.2 − 6.1 T); in the presence of glutamic acid,

stationary phase cells display up to a 100,000 times survival elevation in comparison

to cells maintained in the geomagnetic field. Concurrently, strong fields also cause an


increase in expression of the sigma factor, Sigma S (rpoS ) [160]. As glutamic acid causes

cell death in stationary phase, it appears likely that the magnetic field is modulating

glutamic acid metabolizing enzymes.

7 Effects on gene expression: transcription and translation

AC MFs of moderate flux density (200 − 660 μT, 50 Hz) alter the transcription rate of

the lac operon in E. coli [161]. Sharp “amplitude windows” are observed for this effect,

which are a hint on a non-linear dose dependence. Furthermore while a field strength

of 300 μT suppresses transcription, a field strength of 550 μT results in a substantial

increase. These antagonistic interactions have been attributed to the involvement of

different ions, i.e. Ca2+ and Mg2+ competing for protein-binding sites [162, 163].

AC magnetic fields can induce specific sets of genes. In E. coli an increase in σ32

mRNA (transcription factor) was found for 1.1 mT and 60 Hz [164]. Pulsed square fields

(1.5 mT) elicit an increase in the α subunit of RNA polymerase, and also NusA, in

E. coli. The protein biosynthesis was studied by gel electrophoresis. Thirty proteins were

identified, which were up- or down- regulated by approximately a factor of two [165]. An

important observation in this context is the fact that AC fields can enhance translation,

even in an in vitro system [166]. This shows that the translation machinery itself must be

magnetosensitive, and is not, for example, dependent on the existance of a biomembrane.

Investigations using HeLa cells, though of human origin, generated data that was highly

pertinent to the problem of magnetically induced gene expression. Lin et al. [167] were

able to show that weak, alternating, magnetic fields (8 and 80 μT, 60 Hz) increased the

transcription of mouse or human c-myc genes. This effect was dependent on the presence

of specific electromagnetic response elements located between −353 and −1257 bp relative

to the promoter [168]. Similar response elements were also detected in the promoter region

of the heat shock gene hsp70 [169].

Strong magnetic fields (14.1 T) caused the transcriptional up-regulation of 21 genes,

and the down-regulation of 44 genes, in Shewanella oneidensis ; while at the same time

causing no substantial alterations in growth [136]. No alteration in the profile of stress

proteins occurred after exposing E. coli to AC fields (7.8 − 14 mT, 5 − 100 Hz) [170].

Furthermore, no changes in differential gene expression (microarray analysis) or protein

profile (2-D gel analysis) were obtained with Saccharomyces cerevisiae exposed to AC

magnetic fields (10 − 300 mT, 50 Hz) [171].

In the photosynthetic bacterium, Rhodobacter sphaeroides, magnetic fields of

0.13 − 0.3 T induced a 5-fold increase in porphyrin synthesis, and enhanced expression

of the enzyme 5-aminolevulinic acid dehydratase, which may be caused by elevated gene

expression [172]. AC MFs can modulate the activity of enolase in E. coli ; at 16 Hz a

stimulation is observed, while at 60 Hz a suppression of enolase activity occurs [173].

Propionylcholinesterase activity in the amoebae Dictyostelium was lowered upon expo-

sure to an AC magnetic field of 200 μT and 50 Hz [174]; while at the same time the

fission rate was reduced. This magnetoresponse was, interestingly, adaptive, as it dis-


appeared after a 24-h, lasting exposure. Very strong DC fields (0.13 − 0.3 T) induce, in

Rhodobacter sphaeroides, an increase of 5-aminolevulinic acid dehydratase concentration

predominantly at the magnetic North pole, an effect that was paralleled by increased

porphyrin production [172].

8 Effects on enzyme activity

The fact that MFs can modulate enzyme activities in vitro is a crucial observation, be-

cause it indicates that enzymes may function as magnetoreceptors. Even though several of

the studies listed in Table 6 used enzymes derived from animals or plants, they neverthe-

less show that enzymes have the potential to function as magnetoreceptors. For example,

a static MF of 20 μT alters the in vitro activity of Ca2+/calmodulin-dependent cyclic

nucleotide phosphodiesterase in a Ca2+-dependent manner. This effect shows that the

earth’s magnetic field could be biologically relevant in calcium-dependent reactions [175].

Weak MFs ranging from 0−200 μT modulate the phosphorylation rate of the 20 kDa light

chain of myosin by affecting Ca2+/calmodulin-dependent myosin light chain kinase [176–

179], an observation that remains, however, unconfirmed by other researchers [180]. A

moderate increase or decrease in the geomagnetic field modulates, in vivo and in vitro,

the activity of hydroxyindole-O-methyltransferase (HIOMT, EC 2.1.1.4) and N-acetyl-

serotonin transferase (NAT, EC 2.3.1.5); two key enzymes in the biosynthesis of mela-

tonin in the pineal gland and retina [181]. A 50% increase or decrease in the geomagnetic

field strength caused a decrease of HIOMT activity. NAT responded differently in that

a 50% increase in magnetic field increased activity in the pineal organ, but not in the

retina. The enzyme was unresponsive to a decrease in field strength. These observations

are particularly pertinent in view of a series of investigations on the effects of static and

alternating magnetic fields on human and animal behaviour, and melatonin synthesis.

Numerous studies have shown that magnetic fields can substantially alter circadian mela-

tonin levels [182–185]. One such example is the brook trout (Salvelinus fontinalis), in

which AC-fields (40 mT, 1 Hz) elicit an increased night-time, pineal and serum melatonin

levels [186]. In pigeons the activity of the melatonin-synthesizing enzyme NAT was sub-

stantially reduced in the pineal glands of pigeons exposed, for 30 min at midnight, to a

50 degree rotation in the horizontal component of the earth’s magnetic field [187].

The threshold for stimulation of the Na, K-ATPase by electromagnetic fields is ex-

tremely low, i.e. 0.2 - 0.3 μT [188], a value close to the threshold for transcriptional

stimulation in human cell cultures [189].

Strong MFs (6 T, uniform field) reduced the activity of L-glutamic dehydrogenase by

some 10%, whilst in a non-uniform field of 7 T it was reduced up to 93% [190]. Catalase

was similarly modulated by strong magnetic fields [190]. The activity of carboxydismu-

tase from spinach chloroplasts exposed to a strong magnetic field of 2 T was substantially

enhanced [191]. The activities of trypsin [192] and ornithine decarboxylase [193] can be

enhanced in strong magnetic fields. Weak and moderate, static and alternating, mag-

netic fields (50 Hz) influence the redox activity of cytochrome-C oxidase [194]. Triticum


responds to treatment with 30 mT (50 Hz) with increased esterase activity and proton

extrusion [195]. The activity of horseradish peroxydase (1 mT, 50 − 400 Hz) depends

substantially on the frequency of the applied magnetic field [196].

9 Effects on metabolism

AC fields (14.6 mT, 60 Hz) can provide protection for Salmonella typhimurium from heat

stress [153]. This observation is particularly interesting in view of the fact that magnetic

field exposure can induce the heatshock protein HSP70 in Drosophila [197].

Magnetic fields can exert substantial effects on the metabolic rates of organisms. For

example, Saccharomyces cerevisiae, when exposed to an AC field (0.5 μT, 100− 200 Hz),

responded with a 30% reduction in respiration [198]. Corynebacterium glutamicum in-

creases ATP levels by about 30% in an AC field (4.9 mT, 50 Hz) [199]. In the cyanobac-

terium, Spirulina platensis, a DC field of moderate strength (10 mT) enhanced growth,

O2 evolution, and pigment synthesis; at 70 mT however, a repression, rather than stimula-

tion, was observed [200]. AC fields (0.1 mT, 60 Hz) caused lower ATP levels in Physarum

polycephalum, but no decreased respiration [201]. Reduced respiration was, however,

found with 0.2 mT and 60 and 75 Hz [133]. Tetrahymena pyriformis responds to an AC

field (10 mT, 60 Hz) with delayed cell division and increased oxygen uptake [202].

10 Effects on differentiation: growth patterns and germination

The dimorphic fungus Mycotypha africana can exist in a myceliar or yeast-like form.

Weak ELF magnetic fields shift development towards the yeast form [203]. Weak AC

fields (0 − 1.2 nT, 0.8 − 50 Hz) further increase this germination rate [204]. Very strong

DC fields (5.2−6.1 T) suppress spore formation from vegetative cells of Bacillus subtilis,

an effect that was paralleled with the diminished activity of alkaline phosphatase [135].

11 Effects on behaviour: gravitaxis and bioluminescence

AC fields (0.5 − 2.0 mT, 50 Hz) elicit in the ciliates Paramecium biaurelia, Loxodes

striatus and Tetrahymena thermophila increased swimming velocities and a decrease in

the linearity of cell tracks. At least in the case of Paramecium, this response must be

Ca2+ specific as it is abnormal in Ca2+-channel mutants [205]. Paramecium multimi-

cronucleatum transiently responds to an AC field (600 mT, 60 Hz) with an enhanced

gravitaxis [206]. Nakaoka et al. [207] found no effect of an AC field (650 mT, 60 Hz) on

swimming orientation of Paramecium tetraurelia. The fact that some of these responses

are Ca2+-specific and -dependent is highly relevant in view of the observation that several

magnetic phenomena in animals, such as morphine-induced analgesia in mice [208, 209]

and pineal melatonin synthesis in teleost fish [186], are also related to Ca2+ channels.

Contradictory data exist regarding magnetic effects on bioluminescence. No effect was

found for AC fields for Vibrio fisheri [210], while enhanced bioluminescence was described


for Vibrio quinghaiensis after exposure to AC fields (0.1 − 9.6 mT, 50 Hz) [211]. Geo-

magnetic storms were reported to prolong bioluminescence in Photobacterium [132].

12 Effects on ecology: aquatic systems

The migration and distribution of magnetotactic bacteria in marine and freshwater aquatic

systems is dependent on the magnetization of the local environment [212]. On one hand

this is determined by the petromagnetic properties of benthic deposits, reflecting the

paleo-ecological history of this aquatic biotope [213]. On the other hand, short- and

medium-term variations of the geomagnetic field, e.g. caused by increased solar activ-

ity, are superimposed. A variation of biological productivity, which correlates with the

occurrence of biogenic magnetite, was found in the Rybinsk Reservoir [214]. In the lit-

toral of the same artificial biotope, a correlation between geomagnetic activity, water

transparency and photosynthesis intensity of phytoplankton was investigated [215]. The

ecological role of magnetotactic bacteria in coastal salt ponds, whose spatial and temporal

distribution is affected by the geomagnetic field, is described by Sakaguchi et al. [31].

Liquids, in organisms and natural waters, generally contains colloids, consisting of

dissolved gases, dispersed biological material, small soluble carbonates and other similar

components; all of which contribute to a multitude of boundary layers with concomitant

zeta potentials that originate from these space-charge areas. Exposure of such solutes to

weak MF and EMF caused altered solvation properties for carbonates and gases, and,

in addition, also affected surface tension, viscosity and pH [216, 217]. Furthermore,

the observation that pH alterations in soils, which always contain a wide spectrum of

biocolloids, correlate with geomagnetic events [218] could be explained along these lines.

These findings could be highly relevant for assessing the consequences of the decreases

in main geomagnetic field strength [219] that has been occurring for about 150 years.

This could also likely affect the solubility of CO2, O2, and CH4, as well as the solution

equilibrium of carbonates in the oceans [220–222]. Thus it would be an essential factor

regarding marine carbon cycles.

13 Mechanisms and models for magnetoreception

Models of magnetoreception need to explain sensitivity to: (i) static magnetic fields, (ii)

alternating magnetic fields; and (iii) resolve the paradox that the thermal energy content

of (living) matter exceeds, by many orders of magnitudes, that of the magnetic field.

Chemical reaction rates depend on temperature, and the existence of liquid water, and

occur typically at temperatures of T > 273 K (0◦C), which correspond to a thermal energy

of E > 3.76 ·10−21 J. Weak magnetic fields of far lower energies can nevertheless affect life

processes, which implies that they generate molecular order and overcome the thermal

barrier, i.e. entropy. A theoretical limit for the threshold of a biomagnetic response is

determined by the fact that the magnetic flux is always quantized [223], the magnitude

of the magnetic flux quantum being 2.07 · 10−15 Tm−2. The lowest thresholds that have


been reported for magnetobiological responses approach this value, an observation that

represents a formidable challenge to any theory of magnetoreception [142].

13.1 Ferrimagnetism

Ferrimagnetic magnetoreceptors consist of magnetic minerals like magnetite (Fe3O4) and

greigite (Fe3S4), and act as the magnetoreceptors for bacterial magnetotaxis. In contrast

to ordinary magnets (ferromagnetism), in which the individual magnetic moments of the

electron spins are aligned in parallel, the magnetic moments are antiparallel for ferri-

magnetic materials. Magnetite, which is more precisely written as (Fe3+Fe2+) Fe3+ O2−4 ,

is characterized by a spinell type of crystal structure, i.e. it possesses two unequivalent

lattice positions, tetraedrically coordinated A-positions and octaedrically coordinated

B-positions. The A-positions are occupied exclusively by Fe3+-ions, while the B-positions

are equally occupied by Fe3+- and Fe2+-ions (inverse spinell). In this structure the mag-

netic moments of the Fe3+-ions cancel each other and the residual magnetic moments

derive from the Fe2+-ions. Ferrimagnetism is thus much weaker than that observed for

ferromagnetic materials. On the other hand, the force generated by ferrimagnetic materi-

als exceeds those of dia- or paramagnetic materials by more than 6 orders of magnitude.

Magnetite crystals are organized in magnetosomes, which assemble chains along the

motility axis of the bacterium, generating a permanent magnetic dipole moment and

aligning the cells parallel to the geomagnetic field lines (see above). Magnetites are able

to transport electrons, and thus to conduct current. Whether or not this property plays

a role in biology remains presently unknown. Because magnetite is ubiquitous in the

animal kingdom, it is possible that ferrimagnetic nanocrystals also play a vital role in

organisms other than magnetotactic bacteria. One possibility could be that nanocrystals

fixed to ion channels modulate ion movement across membranes [77]. The energy that

would be absorbed by a single 100-nm magnetite crystal exposed to 0.1 mT at 60 Hz is,

in theory, sufficient to exceed the thermal noise [78].

13.2 Radical-pair mechanism

When two radical molecules stay, for a short time, in close proximity displaying spin cor-

relation, they form a radical pair. This spin correlation of radical pairs implies a normally

forbidden state of equal spin quantum numbers (“parallel spins”, Figure 1), resulting in

a net paramagnetic momentum. There exists a multitude of ways to generate radical

pairs. One possible way is through homolysis of a molecule of 1A-D that is split into two

radicals, A• and D•, creating at first a pair 1[A• D•] in the singlet state (Wigner conser-

vation rule; the single bond in 1A-D also being in a singlet state). Under the influence

of such a magnetic field (including the weak field of a nuclei), the radical pair undergoes

intersystem crossing (ISC) to form a pair 3[A• D•] in the triplet state. Singlet and triplet

radical pairs have different fates: while the singlet pair can recombine directly to the


original donor 1A-D, the triplet cannot and will often take indirect routes. Since a mag-

netic field induces the generation of triplet pairs, the magnetic field effectively causes a

longer lifespan for the radicals. A donor molecule 1A-D may also give rise to a radical

pair made up of cationic and anionic radicals, as shown in Figure 1. Other modes of

radical-pair formation may involve other reaction partners, and redox reactions, as in the

case of cryptochrome (see below). A great advantage of the radical-pair mechanism is

the fact that magnetically modulated ISC and radical pair recombination are inherently

temperature independent, so that no kT -problem arises.

Figure 1 shows an example how homolysis of a donor molecule, 1A-D, leads, via the ex-

cited singlet state 1A-D∗, to the formation of a cation-anion radical pair 1[A•+ D•−]. As a

rule, the mobility of the single radicals - in the specific case A•+ and D•− - is, immediately

after its generation, restricted, because of their size and the viscosity of the environment.

Remaining close together, the partners exist in the singlet state, 1[A•+ D•−], and have

antiparallel spins. ISC provides a paramagnetic state by the spin-orbit coupling of elec-

trons, with the consequence that it can be modulated by an external magnetic field (MF).

This isoenergetic, radiationless transition between two electronic states has different mul-

tiplicities, and enables the interconversion of the pair to the triplet state, 3[A•+ D•−], in

which the single radicals can have parallel and antiparallel spins, implying four possible

states with a probability of 25% each (↑↓, ↓↑, ↑↑, ↓↓). Here, only the hyperfine-niveaus,

with parallel spins, are paramagnetic and can interact with a moderate magnetic flux.

In the presence of a MF (B > 0), the triplet states, with parallel spins T+1, T−1, have a

probability of 50%, and do not contribute to the subsequent reaction.

As long as the external MF is zero or very weak, the three triplet states: T0, T+1,

T−1, of the radical pair 3[A•+ D•−] can recombine to 3AD. In bacterial photosynthetic

reaction centers this allows for the conversion of 3O2 to 1O2 (Figure 1). With increasing

magnetic field strength (B > hyerfine interaction), and concomitant Zeeman splitting,

the probability for recombination of the T+1 or T−1 states decreases, and only the T0

state recombines (Figure 2). For very weak magnetic fields, in the range of the hyperfine

interaction, the yield of the singlet state decreases, while it increases for elevated magnetic

flux densities (Figure 3).

In the framework of the radical-pair mechanism magnetobiological effects are ex-

plained in the following way: (i) external magnetic fields shift the equlibria of singlet and

triplet radical pairs, (ii) the primary magnetobiological response occurs either from the

singlet or else from the triplet radical pair, and (iii) because the fates of the singlet and

the triplet radical pairs are different (Figure 1), it is expected that the requisite biolog-

ical responses are equally different, i.e. dependent on the magnetic flux density. It is

irrelevant in this context whether or not the magnetobiological response occurs from the

singlet or else from the triplet state of the radical pair.


Fig. 1 Generation of cation-anion radical pairs, and ISC (intersystem crossing) under

the influence of a magnetic field. Without a MF (B = 0) the three triplet states may

recombine to the triplet radical 3AD. For B > 0 (Zeeman splitting) only the T0 pairs can

recombine to 3AD, while T+1 and T−1 are excluded from recombination. In photosyn-

thetic reaction centers this leads to the formation of singlet-oxygen (3O2 −→1O2). Other

radical-pair reactions do not, of course, necessarily lead to the generation of 1O2, and the

singlet and triplet radical pairs can have different fates and decay products. Modified

after Liu et al. [237].

Fig. 2 Energy states and spin multiplicities in dependence of the magnetic flux density.

When the energy separation exceeds that of the hyperfine interaction, radicals in the T−1

and T+1 state are excluded from the spin flip, and as a consequence, the singlet yield

increases (Figure 3).


The MF-dependent interconversion rate between the singlet and triplet radical pairs

can be described by the Zeeman interaction. The energy difference ΔE for the two

splitting Zeeman niveaus by a MF (B > 0) is given by the equation:

ΔE = gβB (1)

where g is the Lande factor (near 2 for free electron radicals), B is the magnetic flux

density, and β is the Bohr magneton (9.274 × 10−24 J T−1) [224].

Fig. 3 Dependence of the singlet yield of radical pairs in dependence of the magnetic field.

The magnetic flux density, B, is expressed in relative units as multiples of a, the average

strength of the hyperfine interaction. Modified after Timmel and Henbest [354]. LFE:

low-field effect; ’normal’ MFE: normal magnetic field effects often occur in the mT-range.

Depending on external MF strengths, lifetimes of the occurring spin dynamics (spin

flip) may last up to some μs, and compete with radical separation [225]. ISC is further-

more influenced by hyperfine coupling of the MF with the nucleic magnetic momentum.

Because of the spin relaxation times of about 1 μs, the effects of ELF magnetic fields are

frequency independent up to a few MHz. For a MF of 50 μT to be biologically effective

one requires a cage time of about 50 ns, during which the two partners stay in close

proximity. During this time the hyperfine field of the radicals must allow at least one

precession period; in addition, also the recombination time must be of the same order as

the cage time [226]. Cage times critically depend on the molecular environment; macro-

molecules with cavities or pockets such as nucleic acids or proteins extend the cage times

of radical pairs, and contribute in this way to sensitizing living matter to magnetic fields


of moderate strength [227]. For lower magnetic fluxes, these time windows become still

more critical, so that MFs substantially smaller than the geomagnetic field can hardly

mediate biological effects solely by this mechanism [226]. On the other hand a relief of

MF impacts, by partial thermal decoupling of the electron spins, is suggested by several

authors [226, 228, 229].

In some cases photogenerated radical pairs seem to be involved in magnetoreception,

e.g. for bird orientation [11, 12]. The magnetoreception of migratory birds depends on the

presence of cryptochrome, a FAD-containing, blue-light receptor in the retina [13, 14], that

probably undergoes a photoreduction with concomitant generation of FADH• radicals,

and radical-pair formation. Blue light-mediated, hypocotyl shortening of Arabidopsis

thaliana is clearly influenced by a weak MF of 400 μT [15]. Because double mutants

lacking cryptochromes 1 and 2 do not respond to such fields, this magnetoresponse also

depends on functioning cryptochrome (Figure 4). Because Arabidopsis cryptochrome

generates, upon blue-light absorption, a [FADH• Trp•] radical pair [230], it is very likely

that cryptochrome operates as a magnetoreceptor only in its radical-pair state [15]. In

line with this assumption is the observation that Arabidopsis reacts to a magnetic field

only upon blue-light irradiation, but not, however, in darkness [15].

In the reaction centers of photosynthetic bacteria or plants, absorption of light leads

to the formation of 1Chl, and subsequent charge separation, which includes electron

transfer to a second pigment (PheoChl), thus forming a radical pair in the singlet state,

i.e. 1[Chl+• Pheo−•]. Only relatively strong magnetic fields in the range of several hun-

dred mT can affect the singlet-triplet mixing of the radical pair, because the pair is

rather shortlived. The short half-live of the radical pair is due to the fact that the pair

rapidly donates an electron to a quinone. Technically the described magnetic effects

are monitored by measuring the triplet yield and the fluorescence emission intensity of

the bacterial photosynthetic reaction center [231, 232]. Because the magnetically modu-

lated charge separation processes of bacterial photosynthesis [231, 233], or photosystems

I and II of green plants [234–236], require rather high magnetic flux densities between 10

to several hundred mT, one can conclude that the geomagnetic field does not influence

photosynthetic electron transport.

A very elegant system to study the radical-pair mechanism is a mutant of the purple

bacterium Rhodobacter sphaeroides that lacks carotenoids, so that photosynthetically

generated singlet oxygen is longer lived than in the wild-type strain (1O2 being quenched

by carotenoids). At magnetic flux densities between 0 to 100 mT, the yield of 1O2

decreases in the mutant from 100% to about 50% [237]. The response is predicted by

the model shown in Figure 1. Because of the magnetically-induced Zeeman splitting

for B > 0, only the T0 state is available for recombination and formation of 1O2, while

the triplet states T+1 and T−1 do not contribute (Figure 1). Flash-induced bleaching of

the photosynthetic reaction centers is likewise dependent on magnetic flux densities; for

example, the bleaching at 800 nm at 15 mT is 45% smaller than that in a zero field [237].


Evidence for radical-pair mechanisms were also obtained for some retinoids and por-

phyrins involved in the mitochondrial respiratory chain, where radical pairs could enhance

the synthesis of reactive oxygen species (ROS) [238]. Intermediates of enzyme reactions

may involve the formation of radical pairs. An enzyme that has been investigated in detail

is B12 ethanolamine ammonia lyase [224, 239, 240]. Further cases of radical pair mecha-

nisms include ionizing radiation damage, and its concomitant repair. Dicarlo et al. [241]

report increased repair rates after ultraviolet light (UV) exposure and subsequent treat-

ment with a 60 Hz EMF of only 8 μT field strength, an observation that confirmed earlier

results [242–244]. Magnetic fields also substantially influence antioxidant scavenging of

ROS [245].

Fig. 4 Radical-pair formation of the blue-light receptor cryptochrome upon absorption of

near-UV or blue light. The excited chromophore, S1FAD, undergoes a photoreduction to

form, with a tryptophanyl residue (Trp) from the apoprotein, a radical pair; the electron

and proton donors are omitted. The radical pairs can recombine to form S0FAD, or triplet

products. The biological effector molecule may be derived either from the singlet radical

pair (as shown in the figure) or else from the triplet radical pair (not shown). Modified

after Ahmad et al. [15].

13.3 Ion cyclotron resonance

In the mid eighties it became increasingly clear that numerous biomagnetic responses

display characteristic dose-response relationships, with distinct amplitudes and frequency

“windows”. To explain these results, a physical phenomenon was taken into account, which

was long known from vacuum physics, and thinned gases. Charged particles moving

perpendicularly to a magnetic field are deflected by the Lorentz force on a circular path

perpendicular to the magnetic field lines. An electron orbiting around the nucleus has

a magnetic momentum, which is proportional to its angular momentum, L (Figure 5).


An external MF affects L by an additional torque, ΔL, and forces the electron to precess

around the magnetic field, B0. The resulting precession angular velocity, ωLarmor (Larmor

precession), can be written as:

ωLarmor =dφ

dt=

B0e

2me

(2)

where e is the charge and me, the mass of the electron. There is substantial experimental

evidence that magnetobiological effects are maximal when the frequency of an alternating

MF BAC , which is superposed to a static magnetic field, coincides with the Larmor

frequency of a biological relevant ion such as Ca2+ or Na+. Because of this fundamental

relationship, dose-response curves display the characteristic presence of “windows”.

Fig. 5 The Lamor precession of a charged particle around a magnetic field, B, having

a rotating angular momentum vector, L, that circumscribes the surface of a cone. For

further explanation see text.

The field strength BDC of the MF, the charge Q, and mass m of the involved ion, as

well as the corresponding frequency (f) of the additional superimposed ELF-EMF can

be described by the “ion cyclotron resonance” (ICR) formula”:

f =BDCQ

2πm(3)

Here, the specific charge (Q/m) of ions like Ca2+, K+, Mg2+ is a unique material

constant, determining its circulation frequency on a forced orbit.

One of the first ICR models [246, 247] described Ca2+ ions moving helically along the

geomagnetic field lines. A superposed ELF magnetic field of suitable frequency accelerates

the movement, purportedly resulting in an increase of Ca2+ influx via calcium channels

that are aligned with the geomagnetic field. Biological relevant effects, and in vitro effects,

have been obtained for nearly any ion of characteristic ELF frequency, e.g. [248–250].

Significantly, the globally used powerline frequencies of 50 and 60 Hz are pertinent in this

context [251, 252], as they provide ICR for Ca2+ and many other important ions. Relevant

effects can be obtained, for example, if a 50 Hz AC field of a moderate flux density of 65 μT


superposes with a static field, BDC, that is comparable to the geomagnetic field [253].

One thus has to reckon with the possibility that ICR conditions, for Ca2+ and other ions,

are ubiquitous in our technical environment, with an innate ability to influence health

and biological experiments. Further consequence are found with very weak natural AC

fields caused by the atmospheric Schumann-resonance [254], the circumpolar Birkeland

currents [255] from the auroral zones, and the van Allen radiation belts, each of which

have putatively been persistent ecological and evolutionary factors.

The criticism of earlier ICR models [256, 257] hinges on the problem, that the thermal

energy of biological matter (kT ) is too high, by several orders of magnitude, to allow

the undisturbed movement of charged particles on classical Lorentzian orbits [251, 258,

259]. To resolve this dilemma, theoretical attempts were made to decouple micro-regions,

controlled by weak magnetic fields, from the thermal equilibrium. A possible decoupling

mechanism could consist in a transition zone between molecular layers of decreasing

refraction numbers, for example, water to oil interfaces. All ICR theories imply such

transition zones, typically lipid membranes, tertiary protein structures, cell organells, or

the two-phase state of water (Chapter 13.5 Quantum coherence).

It should be stressed, however, that ICR constitutes a phenomenon that exists even

in the absence of macromolecular structures, such as interface-forming proteins, lipid

membranes or microtubuli, as it occurs even in amino acid solutions exposed to suit-

able combination of BDC and BAC fields [260]. The effect of such fields manifests as

an increase in electric conductivity when the ICR condition for the amino acid is met.

Interestingly, a splitting, in two closely adjacent conductivity bands, became apparent

when the magnitude of BDC was scanned and the conductivity was monitored by an AC

synchronous to the frequency of BAC ; an observation that could indicate a multi-term

energy scheme for the underlying process [261]. These experiments can be understood

in the framework of quantum electrodynamic models, which provide for coherence, and

collision-free movements of small ions [262].

To overcome the kT -problem interactions with electric fields were also taken into

account [263]. It was proposed, for example, that not only a parallel combination of BDC

and BAC , but also a perpendicular arrangement, allows for ICR [264]; an observation

that could indicate rather stable energy states brought about by intrinsic alternating

electric fields, and the geomagnetic field. ICR could be the physical basis for perception

of weak EMF in biological matter, it could furthermore explain substrate specificity,

high sensitivity and unusual dose-response relationships, i.e. “effective windows”, that

are usually absent in other biological dose-response curves. It can also be helpful for

the interpretation of the effects caused by static MF; this would require, however, the

displacement or rotation of charged particles [265], to generate a local BAC. An example

for such a mechanism is the magnetoresponse “anomalous viscosity time dependence” of

E. coli in static magnetic fields (0 − 110 μT). The dose-response curve for this response

shows several prominent maxima and minima (“windows”), which can be explained by

the rotation of ion-protein complexes [19].


13.4 Ion parametric resonance, ion interference mechanism

The original ICR theory of Liboff [246, 247] was later modified by the ion paramag-

netic resonance (IPR) model. The former predicts ELF magnetic effects at the cyclotron

frequencies and their harmonics [266], the latter at the cyclotron frequencies and their

subharmonics [257, 267]. IPR, a generic name for a number of theoretical models based

on classical electrodynamics, as well as quantum electrodynamics, views biomagnetic ef-

fects largely as magnetically modulated ion binding, and thus provides a description for

ion-ligand interactions in MFs. The IPR model also applies to experimental situations in

which a static magnetic field (BDC) is superposed to a parallel ELF magnetic field (BAC);

but theoretical considerations predict low sensitivities in the range of several hundred μT

range [259, 264]. An experimental confirmation of the IPR model was attempted

by Smith et al. [249] with germination experiments, and by Berden et al. [268] using

bioluminescence of the dinoflagellate, Gonyaulax scrippsae.

Ions caged in a protein domain can be described by a superposition of quantum states,

in which the contribution of quantum mechanical interference becomes relevant because

it results in uneven distribution of the ion inside the cage [162, 269]. According to this ion

interference mechanism (IIM), a static magnetic field induces an inhomogeneous density

pattern that begins to rotate with the cyclotron frequency. The addition of an AC MF

results in the cessation of rotation, and finally in the release of the bound ion; a process

that may elicit a biological response. IIM relates the magnetic field parameters to those

of dissociation of ion-protein complexes, and is able to explain a number electromagnetic

phenomena that also include the effect of MF on rotating ion-protein complexes, and the

concomitant dose dependency [19], as well as the effects of pulsed magnetic fields [163].

13.5 Quantum coherence

The quantum coherence mechanism can explain several paradoxical observations, among

which the so-called kT -problem stands most precipitously. The energy content, E, of an

EMF that elicits ICRs is several orders of magnitude smaller than the thermal energy

content (kT ) of the molecules in the EMF. In the case of water, for example, the thermal

energy content at 278.5 K is 1.17 ·109 J m−3, while the magnetic energy content at 0.8 μT

amounts to only 2.6 · 10−7 J m−3. For fermions, i.e. particles, the relation between kT

and magnetic fields can be expressed as:

k · T >> E = B · v · l · Q (4)

where Q is the charge moving along distance l, with speed v, inside a magnetic flux, B.

For bosons, i.e. photons, or likewise ELF-EMF, the relation can be expressed as:

k · T >> E = v · h (5)

where T is the absolute temperature, k the Boltzmann constant, v the frequency and h

the Planck constant. It is a well-known principle in physiology that a stimulus needs to


exceed the kT limit in order to elicit a response. The probability (W ) for overcoming

the Boltzmann-distributed thermal equilibrium at temperature T > 0 K, by an external

energy (E) is approximated by:

W (E) ≈ 1 − e−EkT (6)

In photobiology, for example, the energy content of a single photon surpasses, many

times, kT , thus resulting in a value for W near unity; which in turn means that a response

is elicited. In magnetobiology, however, one encounters quite a different situation. If one

calculates W (E) for T = 293 K, and a MF of B = 40 μT (i.e. geomagnetic field), one

obtains for the spin-related energy of an electron a value for W as low as ∼ 10−7. This

means that statistically only one out of ten million free electrons contributes to a charge

transfer caused by MF interactions. It is apparent that such a low particle fraction could

not possibly elicit a biological reaction. Since weak fields do, however, elicit biological

reactions, it appears on the grounds of eq. 6, as if living matter behaves in a MF like

a subcooled gas at a temperature of ∼ 5 · 10−6 K; which would result in a conduction

band of ≥ 99% occupation, i.e. W (E) is approaching unity. Such behaviour is known for

Bose-Einstein-condensates, which represent matter states near 0 K, and that appear at

first sight to be incompatible with living matter. The coherence mechanism nevertheless

provides the possibility to resolve this paradoxical situation by providing a mechanism

by which the magnetic field action is thermally decoupled from its environment.

Particle properties (e.g. the spin) are described by quantum mechanics by wavefunc-

tions, which express the probability for a certain quantum state with respect to location

and time. If several similar particles (e.g. photons or electrons), are related to the same

wavefunction in a fixed phase ratio (by analogy to synchronously swinging pendulums)

their state is said to represent “coherence”. The De Broglie equation describes the wave-

length λ of such a matter-wave for a fermion particle (e.g. an electron). Because the

probability of location of the particle inside a distance (l) must be 1, it is the mini-

mum coherence length (zero order) for the particle at the same time. With the matter

wavelength λ ≥ l, particle mass m, and kinetic energy E, it will be written as:

λ =h√

E · 2m (7)

If the location probabilities of at least two particles are coherently superimposed, one

obtains one common wavefunction, i.e. a loss of the individual (e.g. fermion) proper-

ties which leads to Schrodinger’s “entanglement” [270]. In this way a newly condensed

matter state originates in which individual particles appear to be “glued” together by

transposition forces like phonons or solitons (Cooper pairs in superconductivity theory).

At this point of the coherence mechanism, the universal mediator of biological pro-

cesses, water, comes into play. Water dipoles assemble spontaneously into self-organizing,

ordered clusters. Quantum electrodynamics predicts two-state aggregates for water con-

sisting of: (i) a bulk phase, which is determined by the thermal equilibrium (water in

Brownian motion), and (ii) ordered clusters (Figures 6–8) [262, 271, 272]. Clusters orig-


inate spontaneously by an in-phase propagation of the oscillating quantum field of de-

localized orbitals of water molecules [273]. Ordered water clusters generate a 12.06 eV

(resp. 2.9 · 1015 Hz) (Figure 6) superradiation-transition [274, 275], which corresponds to

an electromagnetic wavelength of approximately 100 nm, and a quantum coherence over

∼ 1.36 · 107 water molecules which is likely mediated by excitons [272]. These “coher-

ence domains” (CD) should be spheres (Figures 6–8) with a uniform, zero-dimensional

wavefunction, i.e. they should be quantum dots. Seen from the surrounding environment

the CD appears as supramolecular structure with a molecular mass of 217.6 MDa; the

inside is inaccessible to ions except the hydronium-ion (H3O+), considered by energetic

relations. The oscillation strength is expected to reach 8.4·1010 V m−1 at the center [275],

a value which is two orders of magnitude in excess of that of biological membranes; as-

suming thereby a potential of 100 mV across the lipid bilayer. The spherical interface

region (Figures 6, 8) explains the sudden drop of dielectricity from the interior of the CD

(εr ∼ 160) to the incoherent domain (εr < 80). This results in a mean in these values

that closely mirrors that found experimentally for water (εr ∼ 80).

Fig. 6 Model of a water phase organized in spherical coherence domains (CD). Delocalized

molecule orbitals generate a radially oscillating quantum field with 12 eV, which leads to

a 100 nm spherical superradiation with an amplitude of 8.4 · 1010 V m−1 in the center.

Further explanations in the text.

The 2−4 nm thick interface region of the CD (Figures 6–8) appears in many respects

akin to the water transition zone of lipid membranes. A potential trough of about 0.26 eV,

which is predicted by the Born-equation, constitutes a circular ion trap that is responsible

for the experimentally-observed, ICR effects in water. These ions are assembled in this

interface in a plane perpendicular to MF lines, up to a critical density that is reached when


their Debye-Huckel radii interact (Figure 8, left). This “ring” of ions, around the water

CD, represents a one-dimensional coherent particle system, in effect a quantum wire. For

glutamate solutions at 293 K such a quantum wire is composed of approximately 330

glutamate anions [260, 272]. Ionic current measurements have shown that about 36% of

the glutamate anions are organized in this coherent state. The Eigenvalue of the water

CD (eq. 8) matches the total rotation energy of the ion-ring mediated by the BDC . Such a

coupling of a bosonic (CD) and fermionic (ions) quantum system is known as Freshbach-

resonance, which can be described by the Hartree-Fock-Bogoliubov equations [276, 277].

Fig. 7 Schematic diagram of the coupling of a ring of ions to a water coherence domain

(CD): The zero dimensional water CD is a quantum dot and represents a Bose-Einstein-

condensate; the ions behave like a Luttinger liquid, settling around as (1-dimensional)

quantum wire, in a plain perpendicular to the direction of an external MF. Possible quan-

tum statistics, and exchange forces of the states, are demarcated below the pictogram.

Figure 7 sketches the potential interaction at the frontier region of the water CD, lead-

ing to a Freshbach-resonance. The potential trough that generates the ion ring (quantum

wire) surrounds the CD in its immediate vicinity, without any discontinuity or jump. As

a result the trap is constructed by the ions itself; the polarity of the potential depends on

the sign of the ion charge. This potential will be increasingly influenced by thermic noise

in the transition zone to the outer environment. The coherence mechanism also allows for

an alternative structure of the “quantum wire” as there is no absolute requirement for the

synchronous circulation of each ion. Alternatively, an equivalent effect could be achieved

by impulse-perturbation that would circulate along the ion-ring with the ICR frequency,

e.g. mediated by phonons. The ion-ring could also be stabilized by an equivalent number

of counterions (e.g. H3O+), which concentrate in the same plane outside in the surround-

ing, incoherent environment, comparable with the Stern-layer of lipid membranes. The

coherence condition inside the quantum wire itself will be given by the Debye-Huckel radii


ri (h = Planck constant, mion = ion mass, Echem = electrochemical energy given by the

potential, ε = εr · ε0 = permitivity of the solvent (water), NA = Avogadro constant, the

ion strength∏

is given by the ion-concentration ci and -charge zi):

h√2 · mion · Echem

∼√

ε · k · T2 · NA · e2 · ∏ ion - strenght

∏=

1

2

∑i

ciz2i (8)

With glutamate anions one obtains a radius ri of 1.04 nm using the conditions in the

in vitro ICR experiments of Zhadin et al. [260, 261]. How does the coherence mechanism

account for the observed biological effects of alternating MFs? Ions can access the inter-

face region of the water clusters (Figures 6, 8) and orbit (orbit frequency ω) as long as

their Lorentz-radius, specified by the external magnetic flux density BDC, matches (Fig-

ure 8, left). The superimposed, though substantially weaker, EMF BAC with frequency

ω, interferes constructively for ω = Ω, or one of its harmonics (Figure 8, middle). This

way it causes an interfering distortion of the trap geometry, which becomes time-invariant

for the ICR frequency ω and its harmonics n, increasing the probability for decoherence

(Figure 8. center). Such a mechanism could explain the ICR effect for EMF-amplitudes

BAC , down to some nT. An additional electric field, E, amplifies this effect (Figure 8,

right), which causes transitions of the ions to the incoherent environment (Figure 8,

right). Such effects can be measured electrochemically by increases in the ionic current

through the electrolyte solution [261]. The release of ions to the incoherent environment

could elicit subsequent biological reactions. The coherence mechanism provides a good

approximation for the results of in vitro ICR experiments [256, 260–262].

Fig. 8 The coherent-domain model of water. BDC = static magnetic field, BAC = a much

weaker, superimposed electromagnetic field with frequency ω. Ω is the orbit frequency

of the ion in the frontier region of the coherent region. Left: ICR in the undisturbed

circular ion trap without a superimposed field BAC . Center: a superimposed field BAC

with frequency distorts the circular ion trap, so that ions can“break out”to the incoherent

water phase. Right: complete decoherence and emptying of the ion trap by an additional

local electric field E. Modified after [262]; further explanations in the text.


There remain open questions with respect to the fate of the CDs in ultra pure water,

and in the absence of any magnetic field. Also in pure water at room temperature, some

water molecules are dissociated by auto-protolysis, known as the “ion product of water”.

It is possible that these products serve as ions in order to sustain the Freshbach resonance

mechanism of the CDs. At a zero MF, the ICR frequency likewise reaches zero. Even

though a description for this case is not presently available, it is quite possible that CDs

cannot exist in the absence of a MF.

In addition to the coherent “water spheres” described above, cellular macromolecules

were also considered as coherence mediators for biological EMF effects. For example,

microtubules [278], as well as DNA [279], may function as one-dimensional, anisotropic

“quantum wires”. One should, however, keep in mind that the assumptions of these au-

thors were too specific to derive an ICR model of general applicability for living matter

and aqueous solutions. An ICR effect could also be possible at the boundary layers of

collodial solved particles. If the Lamor radii would approach the particle size, the Lorentz

forces would become parallel to the boundary charge layer, forcing it by the additional

magnetic pressure. Findings for the disappearance of the ICR effect in thoroughly de-

gassed, and ultrafiltrated solutions [280], suggest the existence of such a mechanism.

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mT

grow

thin

hibi

tion

,red

uced

viru

lenc

e[3

05]

Serr

atia

mar

cesc

ens

1.49

T,i

nhom

.fie

ldin

hibi

tion

and

stim

ulat

ion

ofgr

owth

[306

]Sh

ewan

ella

onei

dens

is14

.1T

21up

-,44

dow

nreg

ulat

edge

nes,

nogr

owth

effec

t[1

36]

Spir

ulin

apl

aten

sis

10m

Ten

hanc

edgr

owth

,O2

evol

utio

n,pi

gmen

ts[2

00]

70m

Tde

crea

sed

grow

th,O

2ev

olut

ion,

pigm

ents

[200

]St

aphy

loco

ccus

albu

s0.

1−

1m

Tst

imul

atio

nof

grow

than

dm

etab

olis

m[1

37]

Stap

hyloco

ccus

aure

usM

Fof

hum

anbo

dych

ange

dpr

oper

ties

ofliq

uid

wat

er[1

31]

Stap

hyloco

ccus

aure

us5.

08m

Tde

crea

seof

colo

nysi

zean

dnu

mbe

r[3

07]

Stap

hyloco

ccus

aure

us30

−10

0m

Tgr

owth

inhi

bition

unde

rae

robi

osis

[292

]St

aphy

loco

ccus

aure

us30

−10

0m

Tgr

owth

stim

ulat

ion

unde

ran

aero

bios

is[2

92]

Stap

hyloco

ccus

aure

us0.

5−

4T

noeff

ects

ongr

owth

&an

tibi

otic

.se

nsitiv

ity

[295

]St

aphy

loco

ccus

aure

us1.

49T

,inh

om.

MF

inhi

bition

and

stim

ulat

ion

ofgr

owth

[306

]St

aphy

loco

ccus

aure

us1.

49T

grow

thin

hibi

tion

,exp

osur

e-tim

ede

pend

ent

[296

]St

rept

ococ

cus

mut

ants

30−

100

mT

aero

bios

is=

grow

thin

hibi

tion

[292

]un

aero

bios

is=

grow

thst

imul

atio

nSt

rept

omyc

escl

avifor

me

60−

70m

Tal

tere

dco

rem

iafo

rmat

ion

and

rhyt

hm[3

08]

Stre

ptom

yces

mar

inen

sis

3−

15m

Tin

crea

seof

neom

ycin

synt

hesi

s[3

09]

bact

eria

from

Bra

zile

anla

goon

25−

930

μT

mag

neto

taxi

sca

used

bym

agne

toso

mes

[58,

310,

311]

bact

eria

hom

ogen

eous

MF

grow

th,s

hape

,col

ony

size

unal

tere

d[3

12]

bact

eria

1.5

Tgr

owth

inhi

bition

[306

]m

ultice

llula

rpr

okar

yote

high

erth

ange

o-co

mpl

exsw

imm

ing

patt

ern;

sugg

esting

[61]

mag

netic

field

mag

neto

rece

ptio

n,no

tto

rque

slud

gem

icro

bes

80−

300

mT

enha

nced

sedi

men

tation

ofac

tiva

ted

slud

ge[3

13]

was

tew

ater

mic

robe

s0.

35−

0.63

Ten

hanc

edox

idat

ion

ofph

enol

[314

]

Table

2co

nti

nued

Effec

tof

stat

icm

agnet

icfiel

ds

onbac

teri

a.


mag

netic

Org

anis

mflu

xde

nsity

resp

onse

refe

renc

e

Asp

ergi

llus

giga

nteu

sm

utal

ba15

0m

Tre

duct

ion

ofm

ycel

iar

mas

s[3

15]

Asp

ergi

llus

puni

ceus

200

mT

mor

phol

ogic

alch

ange

sin

coni

dia

[316

]A

sper

gillu

sni

ger

200

mT

colo

nypi

gmen

tation

[316

]A

sper

gillu

sni

ger

0.1−

1m

Tst

imul

atio

nof

grow

than

dm

etab

olis

m[3

17]

Alte

rnar

iaal

tern

ata

200

mT

mor

phol

ogic

alch

ange

sin

coni

dia

[316

]A

ltern

aria

alte

rnat

a0.

1−

1m

Tgr

owth

inhi

bition

[314

]A

ltern

aria

alte

rnat

a0.

1−

1m

Tpr

omot

ion

ofco

nidi

afo

rmat

ion

[134

]A

ltern

aria

alte

rnat

a0.

1,0.

5,1

mT

grow

thin

hibi

tion

[140

]A

ltern

aria

alte

rnat

a0.

1,0.

5,1

mT

prom

otio

nof

coni

dia

form

atio

n[1

40]

Can

dida

15m

Tgr

owth

stim

ulat

ion

[144

]Can

dida

30−

60m

Tgr

owth

inhi

bition

[144

]Cur

vula

ria

inae

qual

is0.

1,0.

5,1

mT

grow

thin

hibi

tion

[140

]Cur

vula

ria

inae

qual

is0.

1,0.

5,1

mT

prom

otio

nof

coni

dia

form

atio

n[1

40]

Cur

vula

ria

inae

qual

is0.

1−

1m

Tpr

omot

ion

ofco

nidi

afo

rmat

ion

[134

]Fu

sari

umox

yspo

rum

0.1−

1m

Tin

hibi

tion

ofco

nidi

afo

rmat

ion

[134

,311

]Fu

sari

umcu

lmor

um0.

3T

inhi

bition

ofm

ycel

ialgr

owth

[317

]re

duce

dvi

abili

tyan

dco

nidi

age

rmin

atio

nPen

icilliu

mcl

avifor

me

60−

70m

Tal

tere

dco

rem

iafo

rmat

ion

and

rhyt

hm[3

08]

Sacc

haro

myc

esce

revi

siae

400

μT

30%

inhi

bition

ofbu

dfo

rmat

ion

[139

]Sa

ccha

rom

yces

cere

visiae

460

mT

grow

thin

hibi

tion

[318

]

Table

3E

ffec

tof

stat

icm

agnet

icfiel

ds

onfu

ngi

and

pro

tist

s.


mag

netic

Org

anis

mflu

xde

nsity

resp

onse

refe

renc

e

Sacc

haro

myc

esce

revi

siae

0.5−

0.8

Tst

imul

atio

nof

grow

than

dre

spir

atio

n[3

03]

Sacc

haro

myc

esce

revi

siae

1.5

Tno

effec

ton

grow

th[3

19]

Sacc

haro

myc

esce

revi

siae

0.35

,2.

45m

Tno

effec

ton

grow

th[3

20]

Sacc

haro

myc

esce

revi

siae

hom

ogen

eous

mod

ifica

tion

ofra

diat

ion

dam

age

[242

]Sa

ccha

rom

yces

cere

visiae

7.28

Tpr

e-ex

posu

re:

incr

ease

dU

V-s

urvi

valr

ate

[321

]Sa

ccha

rom

yces

cere

visiae

7.28

Tpo

st-e

xpos

ure:

decr

ease

dU

V-s

urvi

valr

ate

[321

]ye

asts

,m

olds

mag

net

grow

thin

hibi

tion

[312

]Aca

ntha

moe

ba,3

spec

ies

71,10

6m

T14

−71

%de

crea

seof

grow

th[1

38]

Col

pidi

umco

lpod

a50

0−

800

mT

mov

emen

tan

dgr

owth

inhi

bition

[322

]Lo

xoph

yllu

m50

0−

800

mT

mov

emen

tan

dgr

owth

inhi

bition

[322

]Par

amec

ium

<10

0ηT

grow

thac

cele

ration

[323

]Par

amec

ium

enha

nced

leth

ality

inpr

esen

ceof

dyes

[324

]Par

amec

ium

126

mT

redu

ced

velo

city

,di

sorg

aniz

edm

ovem

ents

[325

]Par

amec

ium

tetrau

relia

680

mT

mag

neto

taxi

spe

rpen

dicu

lar

tofie

ldlin

es[2

07]

P.m

ultim

icro

nucl

eatu

m68

0m

Tdi

amag

netic

anisot

ropy

ofci

liaPar

amec

ium

caud

atum

field

grad

ient

4.3

T/m

10−

15%

decr

ease

inpo

pula

tion

[326

]Par

amec

ium

mag

net

wea

kho

rizo

ntal

mag

netic

field

[327

]Par

amec

ium

caud

atum

15.9

,1.

9m

T,m

agne

tav

oida

nce

ofth

eno

rth

pole

[328

]Par

amec

ium

caud

atum

>3

Tal

ignm

ent

with

field

lines

,di

amag

netism

[329

]Sp

iros

tom

umam

bigu

um12

.5T

less

tole

ranc

eof

2,2’

-dip

yrid

yldi

sulfi

de[3

30]

Tri

chom

onas

vagi

nalis

46,1

20m

Tgr

owth

stim

ulat

ion

[331

]Tri

chom

onas

vagi

nalis

220,

320,

420

mT

grow

thin

hibi

tion

[331

]

Table

3co

nti

nued

Effec

tof

stat

icm

agnet

icfiel

ds

onfu

ngi

and

pro

tist

s.


mag

netic

Org

anis

mflu

xde

nsity

resp

onse

refe

renc

e

Bac

illu

ssu

btili

s0.

8,2.

5m

T,0

.8,1

kHz

alte

red

grow

thpa

tter

n,in

crea

sed

grow

th[1

43]

Bac

illu

ssu

btili

s5−

90m

T,0

−0.

3H

zgr

owth

inhi

bition

orst

imul

atio

n[1

44]

Cor

yneb

acte

rium

glut

amic

um4.

9m

T,5

0H

z30

%in

crea

seof

AT

Ple

vel

[199

]Esc

heri

chia

coli

65,97

ηT

,16,

60H

zm

odul

atio

nof

enol

ase

activi

ty[1

73]

Esc

heri

chia

coli

21μT

,2−

24H

zal

tere

dD

NA

-pro

tein

com

plex

es[3

32]

Esc

heri

chia

coli

30μT

,9H

zal

tere

dD

NA

-pro

tein

com

plex

es[3

33]

Esc

heri

chia

coli

0.1−

1m

T,5

0H

zre

duce

dtr

ansp

ositio

nac

tivi

tyof

Tn1

0[1

45]

enha

nced

viab

ility

Esc

heri

chia

coli

1.1

mT

,60

Hz

incr

ease

ofσ

32

mR

NA

,tr

ansc

ript

ion

fact

or[1

64]

Esc

heri

chia

coli

0.07

-1.

1m

T,7

2H

zen

hanc

edtr

ansl

atio

nin

cell-

free

syst

em[1

66]

Esc

heri

chia

coli

1.5

mT

,pul

sed

squa

rem

ore

αsu

buni

tR

NA

poly

mer

ase,

Nus

A[1

65]

Esc

heri

chia

coli

0−

22m

T,1

6&

50H

zsh

orte

ned

gene

ration

tim

e[1

42]

Esc

heri

chia

coli

1−

10m

T,2

−50

Hz

noeff

ect

onpr

otei

nsy

nthe

sis

and

grow

th[3

34]

Esc

heri

chia

coli

0.2−

0.66

mT

,50

Hz

alte

red

synt

hesi

sof

β-g

alac

tosi

dase

[161

]Esc

heri

chia

coli

0.1−

1m

T,5

0H

zre

duct

ion

ofT

n10

tran

spos

itio

nac

tivi

ty[1

55]

sinu

soid

alno

effec

ton

grow

thEsc

heri

chia

coli

0.05

−1

mT

,50

Hz

enha

nced

Tn

10tr

ansp

ositio

n[1

45]

puls

edsq

uare

wav

eno

effec

ton

grow

thEsc

heri

chia

coli

1.2

mT

,50

Hz

enha

nced

Tn5

tran

spos

itio

nby

Dna

K/J

[157

]Esc

heri

chia

coli

0.4−

1.2

mT

,50

Hz

enha

nced

DN

Are

pair

,Dna

K/J

synt

hesi

s[1

56]

Esc

heri

chia

coli

7.8−

14m

T,5

−10

0H

zno

alte

ration

ofst

ress

prot

eins

[170

]Esc

heri

chia

coli

10m

T,5

0H

zde

crea

sed

viab

ility

[146

]Esc

heri

chia

coli

150

mT

,50

Hz

cell

killi

ng,flo

wth

roug

hm

agne

tic

field

[335

]Esc

heri

chia

coli

160

mT

,62

kHz

decr

ease

dsu

rviv

al[1

52]

Esc

heri

chia

coli

wea

kfie

ld,l

owH

zge

nera

tes

part

ialdi

ploi

dsdu

ring

conj

ugat

ion

[336

]aff

ects

reco

mbi

nation

and

grow

thEsc

heri

chia

coli

ELF

grow

thst

imul

atio

n[2

0]

Table

4E

ffec

tof

alte

rnat

ing

mag

net

icfiel

ds

onbac

teri

aan

dbac

teri

ophag

es.


mag

netic

Org

anis

mflu

xde

nsity

resp

onse

refe

renc

e

Fla

voba

cter

ium

spec

.0.

1μT−

4μT

,1,10

Hz

enha

nced

grow

th,c

hang

edm

etab

olis

m[3

37]

Hal

obac

teri

umha

lobi

um5−

90m

T,0

−0.

3H

zgr

owth

inhi

bition

orst

imul

atio

n[1

44]

Lact

obac

teri

umac

idop

hilu

m1

Hz,

10H

zst

imul

ated

grow

th[3

38]

Lis

teri

ain

nocu

a30

−50

kV/m

inac

tiva

tion

,sk

imm

ilk[3

39]

Lecl

erci

aad

enoc

arbo

xyla

ta10

mT

,50

Hz

decr

ease

dvi

abili

ty[1

46]

Pho

toba

cter

ium

phos

phor

icum

1−

10m

T,2

−50

Hz

noeff

ect

onpr

otei

nsy

nthe

sis,

grow

th[3

34]

and

biol

umin

esce

nce

Pro

pion

ibac

teri

umac

nes

200μ

T,50

Hz

noeff

ect

onin

tern

alC

a2+

and

viab

ility

[340

]Pro

teus

vulg

aris

1−

10m

T,2

−50

Hz

noeff

ect

onpr

otei

nsy

nthe

sis

and

grow

th[3

34]

Pse

udom

onas

aero

gino

sa5−

90m

T,0

−0.

3H

zgr

owth

inhi

bition

orst

imul

atio

n[1

44]

Salm

onel

laty

phim

uriu

m0.

2m

T,6

0H

zin

crea

seof

azid

e-in

duce

dre

vert

ants

[158

]Sa

lmon

ella

typh

imur

ium

14.6

mT

,60

Hz

prot

ection

from

heat

stre

ss[1

53]

Salm

onel

laty

phim

uriu

m5−

90m

T,0

−0.

3H

zgr

owth

inhi

bition

orst

imul

atio

n[1

44]

Salm

onel

laty

phim

uriu

m6.

3T

,0.5

Hz

nom

utag

enic

effec

t[3

41]

Salm

onel

laty

phim

uriu

m27

.12

MH

z;2.

45G

Hz

stim

ulat

ion

ofgr

owth

[342

]Se

rrat

iam

arce

scen

s8

mT

grow

thin

hibi

tion

,red

uced

viru

lenc

e[3

05]

Stap

hyloco

ccus

aure

us10

mT

,50

Hz

decr

ease

dvi

abili

ty[1

46]

Stap

hyloco

ccus

epid

erm

idis

5−

90m

T,0

−0.

3H

zgr

owth

inhi

bition

orst

imul

atio

n[1

44]

Vib

rio

fisch

eri

1.3

mT

,60

Hz

noeff

ect

onbi

olum

ines

cenc

e[2

1]V

ibri

oqi

ngha

iens

is0.

1−

9.6

mT

,50

Hz

enha

nced

lum

ines

cenc

ein

’dos

ew

indo

ws’

[21]

RN

A-p

hage

MS2

0.5

mT

,60

Hz

dela

yin

phag

eyi

eld

[343

]ho

st:

Esc

heri

chia

coli

2.5

mT

,60

Hz

impe

ding

repl

icat

ion,

incr

ease

dyi

eld

Table

4co

nti

nued

Effec

tof

alte

rnat

ing

mag

net

icfiel

ds

onbac

teri

aan

dbac

teri

ophag

es.


mag

netic

Org

anis

mflu

xde

nsity

resp

onse

refe

renc

e

Can

dida

albi

cans

5−

90m

T,0

−0.

3H

zgr

owth

inhi

bition

orst

imul

atio

n[1

44]

Myc

otyp

haaf

rica

naw

eak

field

,ELF

mor

eye

ast-

like

form

,bet

ter

germ

inat

ion

[203

]M

ycot

ypha

afri

cana

0−

1.2

nT,0

.8−

50H

zin

crea

sein

germ

inat

ion

[20]

Pis

olithu

stinc

tori

us0.

025,

0.1

mT

,50

Hz

stim

ulat

ion

ofgr

owth

and

ergo

ster

ol[3

44]

Sacc

haro

myc

esce

revi

siae

0.5

μT

,100

−20

0H

z30

%de

pres

sion

ofre

spir

atio

n[1

98]

Sacc

haro

myc

esce

revi

siae

120

μT

,50

Hz

redu

ced

surv

ival

afte

rU

Vir

radi

atio

n[1

59]

Sacc

haro

myc

esce

revi

siae

1m

T,6

0H

zno

addi

tion

alm

utat

ion

rate

s[3

45]

Sacc

haro

myc

esce

revi

siae

0.35

,2.

45m

T,5

0H

zno

effec

ton

grow

th[3

20]

Sacc

haro

myc

esce

revi

siae

10−

300

mT

,50

Hz

nodi

ffere

ntia

lgen

eex

pres

sion

[171

]Sa

ccha

rom

yces

cere

visiae

stim

ulat

ion

ofre

spir

atio

n[3

46]

Sacc

haro

myc

esce

revi

siae

2−

620

μT

,100

kHz

upto

30%

grow

thst

imul

atio

n[3

47]

Sacc

haro

myc

esce

revi

siae

0.11

mV

/m,8

0H

zst

imul

ated

CO

2pr

oduc

tion

[348

]Sc

lero

tium

rolfs

ii0.

5−

20H

zre

duct

ion

ofgr

owth

and

germ

inat

ion

[349

]

Table

5E

ffec

tof

alte

rnat

ing

mag

net

icfiel

ds

onfu

ngi

and

pro

tist

s.


mag

netic

Org

anis

mflu

xde

nsity

resp

onse

refe

renc

e

yeas

t,co

ld-s

tres

sed

ferm

enta

tion

[348

]D

icty

oste

lium

disc

oide

um20

0μT

,50

Hz

decr

ease

offis

sion

rate

,m

odul

atio

nof

prop

iony

lcho

lines

tera

seac

tivi

ty[1

74]

Dic

tyos

teliu

mdi

scoi

deum

0.4

mT

,tra

ins

of2

ms

dam

ping

ofad

enin

enu

cleo

tide

osci

llation

s[3

50]

puls

esga

ted

at20

ms

chan

ges

inph

ase

rela

tion

ship

Gon

yaul

axsc

ripp

sae

1.2,

11.5

mT

,50

Hz

biol

umin

esce

nce

[268

]Lo

xode

sst

riat

us0.

5−

2.0

mT

,50

Hz

incr

ease

dsw

imm

ing

velo

city

[250

]Par

amec

ium

biau

relia

0.5−

2.0

mT

,50

Hz

incr

ease

dsw

imm

ing

velo

city

,[2

05]

abno

rmal

resp

onse

inC

a2+-c

hann

elm

utan

tP.m

ultim

icro

nucl

eatu

m60

0m

T,6

0H

zen

hanc

edgr

avitax

is,tr

ansi

ent

resp

onse

[206

]Par

amec

ium

tetrau

relia

1.8

mT

,72

Hz

incr

ease

dce

lldi

visi

onra

te,C

a2+-s

peci

fic[1

47]

decr

ease

dm

embr

ane

fluid

ity

Par

amec

ium

tetrau

relia

650

mT

,60

Hz

noeff

ect

onsw

imm

ing

orie

ntat

ion

[207

]Phy

saru

mpo

lyce

phal

um0.

2m

T,6

0,75

Hz

dela

yof

mitot

iccy

cle

[148

]Phy

saru

mpo

lyce

phal

um0.

2m

T,6

0,75

Hz

mitot

icde

lay;

decr

ease

dre

spir

atio

n[1

33]

Phy

saru

mpo

lyce

phal

um0.

1m

T,6

0H

zlo

wer

AT

Ple

vel;

node

crea

sed

resp

irat

ion

[201

]Phy

saru

mpo

lyce

phal

um0.

2m

T,7

5H

zin

crea

seof

mitot

iccy

cle

leng

th[1

51]

Phy

saru

mpo

lyce

phal

um0.

2m

T,7

5H

zin

crea

seof

mitot

iccy

cle

leng

th[1

49,1

50]

Tet

rahy

men

ath

erm

ophi

la0.

5−

2.0

mT

,50

Hz

incr

ease

dsw

imm

ing

velo

city

[205

]Tet

rahy

men

apy

rifo

rmis

10m

T,6

0H

zde

laye

dce

lldi

visi

on,i

ncre

ased

[202

]ox

ygen

upta

ke

Table

5co

nti

nued

Effec

tof

alte

rnat

ing

mag

net

icfiel

ds

onfu

ngi

and

pro

tist

s.


subs

trat

em

agne

tic

enzy

me

flux

dens

ity

resp

onse

refe

renc

e

Na,

K-A

TPas

e0.

2−

2μT

thre

shol

dfo

rst

imul

atio

n[1

88]

cycl

icnu

cleo

tide

20μT

50%

activa

tion

,pu

reen

zym

e[1

75]

phos

phod

iest

eras

eC

a2+

and

calm

odul

inde

pend

ent

hydr

oxyi

ndol

e-O

-met

hyltra

nsfe

rase

∼25

μT

20%

decr

ease

ofac

tivi

ty,cr

ude

extr

act

[181

]N

-ace

tyl-s

erot

onin

tran

sfer

ase

∼25

μT

10%

decr

ease

ofac

tivi

ty,cr

ude

extr

act

[181

]hy

drox

yind

ole-

O-m

ethy

ltra

nsfe

rase

∼70

μT

50%

decr

ease

ofac

tivi

ty,cr

ude

extr

act

[181

]m

yosi

nlig

htch

ain

kina

se0−

200

μT

activa

tion

,C

a2+

and

calm

odul

inde

pend

ent

[177

–179

]m

yosi

nlig

htch

ain

kina

se0−

400

μT

kina

sefr

omch

icke

n,no

effec

t[1

80]

B12

etha

nola

min

e-am

mon

ialy

ase

0.1

Tra

dica

l-pai

rm

echa

nism

;25%

enzy

me

inhi

bition

[239

]tr

ypsi

n0.

5T

activi

tyst

imul

atio

n,pu

reen

zym

e[1

92]

carb

oxyd

ism

utas

e2

T20

%st

imul

atio

n,pu

rifie

den

zym

e[1

91]

cata

lase

6T

stim

ulat

ion

16−

52%

[190

]L-g

luta

mat

ede

hydr

ogen

ase

6T

10%

inhi

bition

ina

unifo

rmfie

ld[1

90]

L-g

luta

mat

ede

hydr

ogen

ase

7T

93%

inhi

bition

ina

non-

unifo

rmfie

ld[1

90]

aden

ylat

eki

nase

ofth

e25

0μT

,75

Hz

bovi

nere

tina

;de

crea

seof

activi

ty[3

51]

rod

oute

rse

gmen

tcy

toch

rom

e-C

oxid

ase

10or

50m

T,5

0H

z90

%ch

ange

ofac

tivi

ty[1

94]

from

beef

hear

t30

0μT

or10

mT

90%

chan

ge,n

och

ange

sat

othe

rfie

lds

[194

]ho

rser

adis

hpe

roxy

dase

1m

T,5

0−

400

Hz

activi

tyis

freq

uenc

yde

pend

ent

[196

]ho

rser

adis

hpe

roxy

dase

0−

0.25

T,s

tatic

field

noeff

ect

[224

]or

nith

ine

deca

rbox

ylas

e5

mT

,60

Hz

50%

stim

ulat

ion

ofac

tivi

ty[1

93]

from

L92

9fib

robl

asts

expo

sure

ofce

llsth

resh

old∼

5μT

resp

irat

ory

enzy

mes

cons

tant

effec

tson

activi

ty[3

52]

enzy

me

kine

tics

radi

calpa

irre

com

bina

tion

[240

]es

tera

ses,

Tri

ticu

m30

mT

,50

Hz

invi

votr

eatm

ent,

incr

ease

dac

tivi

ty[1

95,3

53]

incr

ease

dpr

oton

extr

usio

n

Table

6E

ffec

tsof

stat

ican

dal

tern

atin

gm

agnet

icfiel

ds

onen

zym

es.

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